Seedling performance of Pinus densata and its parental population in the habitat of P. tab- uliformis

doi:10.3724/SP.J.1258.2013.00016

Abstract

Abstract:

Aims Pinus densata is the natural diploid hybrid of P. tabuliformis and P. yunnanensis, and is one of a few known examples of homoploid hybrid speciation. In order to describe the adaptive and growth characteristics among the three pine species at seedling stage and clarify the role of ecological selection in interspecific isolation between hybrid and parental species, we conducted a common garden comparative analysis of fitness traits at seedling stage in this species complex.
Methods We investigated 14 adaptive and growth characters in seedlings from 25 representative populations of the three pine species in a typical habit of P. tabuliformis. The 14 traits are germination rate, seedling survival in Aug., Sept. and Nov. 2011, winter survival rate of one-year old seedlings, seedling survival rate in Oct. 2012, rates of bud-set in Nov. 2011 and Oct. 2012, the proportion of yellow, green and purple seedlings, seedling height, ground diameter and crown height of seedlings. Patterns of the variation of these characters were analyzed using multi-factor nested analysis of variance and correlated to geographical factors of each population.
Important findings Most of the traits were significantly differentiated among species and among populations at seedling stage, and the large amount of variance were partitioned among species or among populations. Germination rate, the rates of bud-set in two years, and the survival rate in Oct. 2012 of two-year old seedlings were the best indicators of among-species divergence. P. tabuliformis showed the highest values in most of the traits and appeared as more fit in its native habits than the other two pines. P. yunnanensis was at disadvantage in several critical traits, e.g. the survival rates of Aug., Sept., Nov. 2011 and rates of bud-set in two years. Most of one-year old seedlings of P. yunnanensis have not bud-set at the end of Nov. and the proportion of yellow seedlings were highest and its survival rate in Oct. 2012 was zero, which illustrates its low fitness in P. tabuliformis habitat. The hybrid pine, P. densata was intermediate between the parental species in most of traits except indice of the proportion of purple seedlings. The populations from the ancestry hybrid zone in the northeast of P. densata range coped with the test site environment better than the populations from the western range. Our study highlights the role of ecological selection in speciation of this homoploid hybrid pine.

null

Key words: growth and development, Pinus densata, Pinus tabuliformis, Pinus yunnanensis, adaptation

LIANG Dong, MAO Jian-Feng, ZHAO Wei, ZHOU Xian-Qing, YUAN Hu-Wei, WANG Li-Ming, XING Fang-Qian, WANG Xiao-Ru, LI Yue. Seedling performance of Pinus densata and its parental population in the habitat of P. tab- uliformis[J]. Chin J Plant Ecol, 2013, 37(2): 150-163.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.3724/SP.J.1258.2013.00016

https://www.plant-ecology.com/EN/Y2013/V37/I2/150

Figures/Tables 6

References 41

[1]	Aronld ML, Hondges SA (1995). Are natural hybrids fit or unfit relative to their parents? Trends in Ecology and Evolution, 10(2), 67-71. DOI URL
[2]	Billings WD, Mooney HA (1968). The ecology of arctic and alpine plants. Biological Reviews, 43, 481-529.
[3]	Buerkle CA, Morris RJ, Asmussen MA, Rieseberg LH (2000). The likelihood of homoploid hybrid speciation. Heredity, 84, 441-451.
[4]	Campbell DR, Waser NM (2007). Evolutionary dynamics of an Ipomopsis hybrid zone: confronting models with lifetime fitness data. The American Naturalist, 169, 298-310.
[5]	Day TA, Howells BW, Ruhland CT (1996). Changes in growth and pigment concentrations with leaf age in pea under modulate UV-B radiation field treatment. Plant, Cell & Environment, 19, 101-108.
[6]	Fu JR (1985). Seed Physiology. Science Press, Beijing. (in Chinese)
	[ 傅家瑞 (1985). 种子生理. 科学出版社, 北京.]
[7]	Gross BL, Rieseberg LH (2005). The ecological genetics of homoploid hybrid speciation. Journal of Heredity, 96, 241-252.
[8]	Grubb PJ (1977). The maintenance of species richness in plant communities: the importance of the regeneration niche. Biology Reviews, 52, 107-145.
[9]	Guan ZT (1981). Fundamental features of the distribution of Coniferae in Sichuan. Acta Phytotaxonomica Sinica, 19, 393-407. (in Chinese with English abstract)
	[ 管中天 (1981). 四川松杉类植物分布的基本特征. 植物分类学报, 19, 393-407.]
[10]	Harper JT, Williams JT, Sagar GR (1965). The behavior of seeds in soil. I. The heterogeneity of soil surfaces and its role in determining the establishment of plants from seed. Journal of Ecology, 53, 273-286.
[11]	Heydecker W (1973). Seed Ecology. Butterworths, London.
[12]	Hobbs RJ, Mooney HA (1985). Community and population dynamics of serpentine grassland annuals in relation to gopher disturbance. Oecologia, 67, 342-351.
[13]	Hurme P, Repo T, Savolainen O, Pääkkönen T (1997). Climatic adaptation of bud set and frost hardiness in Scots pine (Pinus sylvestris). Canadian Journal of Forest Research, 27, 716-723.
[14]	Jackson SD (2009). Plant responses to photoperiod. New Phytologist, 181, 517-531.
[15]	Liu ZL, Zhang D, Hong DY, Wang XR (2003a). Chromosomal localization of 5S and 18S-5. 8S-25S ribosomal DNA sites in five Asian pines using fluorescence in situ hybridization. Theoretical and Applied Genetics, 106, 198-204.
[16]	Liu ZL, Zhang D, Wang XQ, Ma XF, Wang XR (2003b). Intragenomic and interspecific 5S rDNA sequence variation in five Asian pines. American Journal of Botany, 90, 17-24.
[17]	Lu ZY (1983). The relationship between the photoperiod and bud-set of Dahurian Larch (Larix gmelinii Rupr.) seedlings. Scientia Silvae Sinicae, 19, 322-326. (in Chinese)
	[ 卢振元 (1983). 兴安落叶松幼苗早封顶与光周期的关系. 林业科学, 19, 322-326.]
[18]	Ma F, Zhao CM, Milne R, Ji MF, Chen LT, Liu JQ (2010). Enhanced drought-tolerance in the homoploid hybrid species Pinus densata: implication for its habitat divergence from two progentors. New Phytologist, 185, 204-216.
[19]	Mao JF (2008). Ecological Genetics of the Homoploid Hybrid Speciation in Pinus densata. PhD dissertation, Institute of Botany, Chinese Academy of Sciences, Beijing. 40-41. (in Chinese with English abstract)
	[ 毛建丰 (2008). 高山松同倍性杂交物种形成的生态遗传学研究. 博士学位论文, 中国科学院植物研究所, 北京. 40-41.]
[20]	Mao JF, Li Y, Wang XR (2009). Empirical assessment of the reproductive fitness components of the hybrid pine Pinus densata on the Tibetan Plateau. Evolutionary Ecology, 23, 447-462.
[21]	Mao JF, Wang XR (2011). Distinct niche divergence characterizes the homoploid hybrid speciation of Pinus densata on the Tibetan Plateau. The American Naturalist, 177(4), 1-16.
[22]	McCarthy EM, Asmussen MA, Anderson WW (1995). A theoretical assessment of recombinational speciation. Heredity, 74, 502-509.
[23]	Nilsson JE, Walfridsson EA (1995). Phenological variation among plus-tree clones of Pinus sylvestris (L.) in northern Sweden. Silvae Genetica, 44, 20-28.
[24]	Pan HL, Li MH, Cai XH, Wu J, Du Z, Liu XL (2009). Responses of growth and ecophsiology of plants to altitude. Ecology and Environmental Sciences, 18, 722-730. (in Chinese with English abstract)
	[ 潘红丽, 李迈和, 蔡小虎, 吴杰, 杜忠, 刘兴良 (2009). 高海拔梯度上的植物生长与生理生态特性. 生态环境学报, 18, 722-730.]
[25]	Rieseberg LH (1991). Homoploid reticulate evolution in Helianthus (Asteraceae): evidence from ribosomal genes. American Journal of Botany, 78, 1218-1237.
[26]	Rieseberg LH (1997). Hybrid origins of plant species. Annual Review of Ecology and Systematics, 28, 359-389.
[27]	Savolainen O, Bokma F, García-Gil R, Komulainen P, Repo T (2004). Genetic variation in cessation of growth and frost hardiness and consequences for adaptation of Pinus sylvestris to climatic changes. Forest Ecology and Management, 197, 79-89.
[28]	Shi SB, Ben GY, Han F (1998). Analysis of the solar UV-B radiation and plant UV-B-absorbing compounds in different regions. Acta Phytoecological Sinica, 23, 529-535. (in Chinese with English abstract)
	[ 师生波, 贲桂英, 韩发 (1998). 不同海拔地区紫外线B辐射状况及植物叶片紫外线吸收物质含量的分析. 植物生态学报, 23, 529-535.]
[29]	Singh A (1996). Growth, physiological and biochemical responses of three tropical legumes to enhanced UV-B radiation. Canadian Journal of Botany, 74, 135-139.
[30]	Song BH, Wang XQ, Wang XR, Ding KY, Hong DY (2003). Cytoplasmic composition in Pinus densata and population establishment of the diploid hybrid pine. Molecular Ecology, 12, 2995-3001.
[31]	Song BH, Wang XQ, Wang XR, Sun LJ, Hong DY, Peng PH (2002). Maternal lineages of Pinus densata, a diploid hybrid. Molecular Ecology, 11, 1057-1063. DOI URL
[32]	Szmidt AE, Wang XR (1993). Molecular systematics and genetic differentiation of Pinus sylvestris (L.) and P. densiflora (Sieb. Et Zucc.). Theoretical and Applied Genetics, 86, 159-165.
[33]	Wang BS, Mao JF, Gao J, Zhao W, Wang XR (2011). Colonization of the Tibetan Plateau by the homoploid hybrid pine Pinus densata. Molecular Ecology, 20, 3796-3811.
[34]	Wang XR, Szmidt AE (1990). Evolutionary analysis of Pinus densata (Masters), a putative tertiary hybrid. 2. A study using species-specific chloroplast DNA markers. Theoretical and Applied Genetics, 80, 641-647.
[35]	Wang XR, Szmidt AE, Lewandowski A, Wang ZR (1990). Evolutionary analysis of Pinus densata Masters, a putative tertiary hybrid. 1. Allozyme variation. Theoretical and Applied Genetics, 80, 635-640.
[36]	Wang XR, Szmidt AE, Savolainen O (2001). Genetic composition and diploid hybrid speciation of a high mountain pine, Pinus densata, native to the Tibetan Plateau. Genetics, 159, 337-346.
[37]	Woodall GS, Steward GR (1998). Do anthocyanins play a role in UV protection of the red juvenile leaves of Syzygium? Journal of Experimental Botany, 49, 1447-1450.
[38]	Wu ZL (1956). The taxonomic revision and phytogeographical study of Chinese pines. Acta Phytotaxonomica Sinica, 5, 131-163. (in Chinese)
	[ 吴中伦 (1956). 中国松属的分类与分布. 植物分类学报, 5, 131-163.]
[39]	Zhang YM, Li GQ, Li LF, Liu DY, Zhao MC (2010). The effect of temperature and humidity on seed germination of Pinus yunnanensis. Shaanxi Forest Science and Technology, ( 4), 1-4. (in Chinese with English abstract)
	[ 张跃敏, 李根前, 李莲芳, 刘代艺, 赵敏冲 (2010). 温度和湿度对云南松种子发芽的影响. 陕西林业科技, ( 4), 1-4.]
[40]	Zheng WJ (1983). Woody Flora of China Volume I. China Forestry Publishing House, Beijing. (in Chinese)
	[ 郑万钧 (1983). 中国树木志: 第一卷. 中国林业出版社, 北京.]
[41]	Zhongshang B (1988). Germination Physiology. Translated by Ma YB. China Agriculture Press, Beijing. (in Chinese)
	[ Zhongshang B (1988). 发芽生理学. 马云彬, 译. 中国农业出版社, 北京.]

物种 Species	采集地点 Collection site	缩写 Abbreviation	纬度 Latitude	经度 Longitude	海拔 Elevation (m)
油松 P. tabuliformis	a 山西方山 Fangshan, Shanxi	FSPt	37°56′ N	111°33′ E	1 855
	b 青海互助 Huzhu, Qinghai	HZPt	36°58′ N	102°58′ E	2 299
	c 四川九寨沟 Jiuzhaigou, Sichuan	JZGPt	33°18′ N	103°47′ E	2 281
	d 山西灵空山 Lingkongshan, Shanxi	LKSPt	36°37′ N	112°20′ E	1 654
	e 河南卢氏 Lushi, Henan	LSPt	33°44′ N	110°49′ E	1 908
	f 内蒙古宁城 Ningcheng, Nei Mongol	NCPt	42°17′ N	118°58′ E	1 200
	g 陕西宁陕 Ningshan, Shaanxi	NSPt	33°29′ N	108°23′ E	1 423
	h 北京松山 Songshan, Beijing	SSPt	40°31′ N	115°49′ E	803
	i 内蒙古土默特 Tumed, Nei Mongol	TMTPt	40°47′ N	111°13′ E	1 195
	j 河北平泉 Pingquan, Hebei	PQPt	41°00′ N	118°16′ E	526
云南松 P. yunnanensis	k 云南保山 Baoshan, Yunnan	BSPy	24°28′ N	99°08′ E	1 935
	l 云南贡山 Gongshan, Yunnan	GSPy	25°58′ N	98°49′ E	1 689
	m 云南昆明 Kunming, Yunnan	KMPy	24°58′ N	102°38′ E	2 240
	n 云南丽江 Lijiang, Yunnan	LJPy	26°53′ N	100°14′ E	2 449
	o 云南宜良 Yiliang, Yunnan	YLPy	24°43′ N	103°10′ E	1 837
	p 云南玉溪 Yuxi, Yunnan	YXPy	24°15′ N	102°10′ E	1 832
	q 云南中甸 Zhongdian, Yunnan	ZDPy	28°01′ N	99°32′ E	3 054
高山松 P. densata	r 四川宝兴 Baoxing, Sichuan	BXPd	30°49′ N	102°44′ E	2 452
	s 西藏察隅 Zayü, Xizang	CYPd	29°80′ N	97°13′ E	3 261
	t 四川康定 Kangding, Sichuan	KDPd	30°11′ N	101°55′ E	2 900
	u 四川理县 Lixian, Sichuan	LXPd	31°40′ N	102°48′ E	2 762
	v 四川马尔康 Barkam, Sichuan	MKPd	31°55′ N	102°12′ E	2 678
	w 西藏八一 Bayi, Xizang	BYPd	29°44′ N	94°05′ E	3 107
	x 西藏帕隆藏布 Parlungzangbo, Xizang	PLPd	29°55′ N	95°30′ E	2 658
	y 西藏米林县 Mainling, Xizang	MLPd	29°14′ N	94°14′ E	2 966

物种 Species	采集地点 Collection site	缩写 Abbreviation	纬度 Latitude	经度 Longitude	海拔 Elevation (m)
油松 P. tabuliformis	a 山西方山 Fangshan, Shanxi	FSPt	37°56′ N	111°33′ E	1 855
	b 青海互助 Huzhu, Qinghai	HZPt	36°58′ N	102°58′ E	2 299
	c 四川九寨沟 Jiuzhaigou, Sichuan	JZGPt	33°18′ N	103°47′ E	2 281
	d 山西灵空山 Lingkongshan, Shanxi	LKSPt	36°37′ N	112°20′ E	1 654
	e 河南卢氏 Lushi, Henan	LSPt	33°44′ N	110°49′ E	1 908
	f 内蒙古宁城 Ningcheng, Nei Mongol	NCPt	42°17′ N	118°58′ E	1 200
	g 陕西宁陕 Ningshan, Shaanxi	NSPt	33°29′ N	108°23′ E	1 423
	h 北京松山 Songshan, Beijing	SSPt	40°31′ N	115°49′ E	803
	i 内蒙古土默特 Tumed, Nei Mongol	TMTPt	40°47′ N	111°13′ E	1 195
	j 河北平泉 Pingquan, Hebei	PQPt	41°00′ N	118°16′ E	526
云南松 P. yunnanensis	k 云南保山 Baoshan, Yunnan	BSPy	24°28′ N	99°08′ E	1 935
	l 云南贡山 Gongshan, Yunnan	GSPy	25°58′ N	98°49′ E	1 689
	m 云南昆明 Kunming, Yunnan	KMPy	24°58′ N	102°38′ E	2 240
	n 云南丽江 Lijiang, Yunnan	LJPy	26°53′ N	100°14′ E	2 449
	o 云南宜良 Yiliang, Yunnan	YLPy	24°43′ N	103°10′ E	1 837
	p 云南玉溪 Yuxi, Yunnan	YXPy	24°15′ N	102°10′ E	1 832
	q 云南中甸 Zhongdian, Yunnan	ZDPy	28°01′ N	99°32′ E	3 054
高山松 P. densata	r 四川宝兴 Baoxing, Sichuan	BXPd	30°49′ N	102°44′ E	2 452
	s 西藏察隅 Zayü, Xizang	CYPd	29°80′ N	97°13′ E	3 261
	t 四川康定 Kangding, Sichuan	KDPd	30°11′ N	101°55′ E	2 900
	u 四川理县 Lixian, Sichuan	LXPd	31°40′ N	102°48′ E	2 762
	v 四川马尔康 Barkam, Sichuan	MKPd	31°55′ N	102°12′ E	2 678
	w 西藏八一 Bayi, Xizang	BYPd	29°44′ N	94°05′ E	3 107
	x 西藏帕隆藏布 Parlungzangbo, Xizang	PLPd	29°55′ N	95°30′ E	2 658
	y 西藏米林县 Mainling, Xizang	MLPd	29°14′ N	94°14′ E	2 966

性状 Character	物种 Species	平均值 Mean	标准偏差 SD	变异系数 CV (%)
出苗率 Germination rate (%)	油松 Pinus tabuliformis	74.050^b	5.733	7.742
	高山松 P. densata	57.875^a	13.699	23.670
	云南松 P. yunnanensis	54.143^a	6.334	11.700
2011年8月的保存率 Survival rate in Aug. 2011 (%)	油松 P. tabuliformis	91.000^a	7.139	7.845
	高山松 P. densata	85.094^ab	12.249	14.395
	云南松 P. yunnanensis	75.607^b	11.341	15.000
2011年9月的保存率 Survival rate in Sept. 2011 (%)	油松 P. tabuliformis	89.450^b	8.065	9.016
	高山松 P. densata	83.344^ab	11.890	14.266
	云南松 P. yunnanensis	74.179^a	10.906	14.702
2011年11月的保存率 Survival rate in Nov. 2011 (%)	油松 P. tabuliformis	88.425^b	7.729	8.741
	高山松 P. densata	82.438^ab	11.560	14.023
	云南松 P. yunnanensis	73.393^a	10.441	14.226
一年生苗越冬保存率 Winter survival rate of one-year old seedling (%)	油松 P. tabuliformis	84.000^c	7.916	9.424
	高山松 P. densata	76.250^b	10.873	14.260
	云南松 P. yunnanensis	67.714^a	9.087	13.420
2012年10月的保存率 Survival rate in Oct. 2012 (%)	油松 P. tabuliformis	63.900^c	9.769	15.288
	高山松 P. densata	18.125^b	30.381	167.619
	云南松 P. yunnanensis	0.143^a	0.378	264.336
2011年11月的封顶率 Rate of bud-set in Nov. 2011 (%)	油松 P. tabuliformis	91.425^c	4.033	4.411
	高山松 P. densata	70.906^b	14.282	20.142
	云南松 P. yunnanensis	10.928^a	20.478	187.390
2012年10月的封顶率 Rate of bud-set in Oct. 2012 (%)	油松 P. tabuliformis	81.900^c	6.976	8.518
	高山松 P. densata	19.834^b	35.432	178.643
	云南松 P. yunnanensis	0.000^a	0.000	-
黄苗比例 Proportion of yellow seedlings (%)	油松 P. tabuliformis	2.300^a	6.845	297.609
	高山松 P. densata	0.406^a	0.481	118.473
	云南松 P. yunnanensis	43.143^b	37.396	86.679
绿苗比例 Proportion of green seedlings (%)	油松 P. tabuliformis	54.350^c	12.849	23.641
	高山松 P. densata	20.688^b	28.561	138.056
	云南松 P. yunnanensis	8.500^a	10.598	124.682
紫苗比例 Proportion of purple seedlings (%)	油松 P. tabuliformis	43.350^a	13.838	31.922
	高山松 P. densata	78.844^b	28.625	36.306
	云南松 P. yunnanensis	47.429^a	34.942	73.672
苗高 Seedling height (cm)	油松 P. tabuliformis	8.286^a	1.242	14.989
	高山松 P. densata	7.763^b	1.070	13.783
	云南松 P. yunnanensis	6.363^c	1.610	25.303
苗冠高 Crown height of seedling (cm)	油松 P. tabuliformis	6.807^a	1.351	19.847
	高山松 P. densata	6.558^b	0.987	15.050
	云南松 P. yunnanensis	5.140^c	1.493	29.047
地径 Ground diameter (mm)	油松 P. tabuliformis	2.486^a	0.124	4.988
	高山松 P. densata	2.093^b	0.270	12.900
	云南松 P. yunnanensis	2.501^a	0.348	13.914

性状 Character	物种 Species	平均值 Mean	标准偏差 SD	变异系数 CV (%)
出苗率 Germination rate (%)	油松 Pinus tabuliformis	74.050^b	5.733	7.742
	高山松 P. densata	57.875^a	13.699	23.670
	云南松 P. yunnanensis	54.143^a	6.334	11.700
2011年8月的保存率 Survival rate in Aug. 2011 (%)	油松 P. tabuliformis	91.000^a	7.139	7.845
	高山松 P. densata	85.094^ab	12.249	14.395
	云南松 P. yunnanensis	75.607^b	11.341	15.000
2011年9月的保存率 Survival rate in Sept. 2011 (%)	油松 P. tabuliformis	89.450^b	8.065	9.016
	高山松 P. densata	83.344^ab	11.890	14.266
	云南松 P. yunnanensis	74.179^a	10.906	14.702
2011年11月的保存率 Survival rate in Nov. 2011 (%)	油松 P. tabuliformis	88.425^b	7.729	8.741
	高山松 P. densata	82.438^ab	11.560	14.023
	云南松 P. yunnanensis	73.393^a	10.441	14.226
一年生苗越冬保存率 Winter survival rate of one-year old seedling (%)	油松 P. tabuliformis	84.000^c	7.916	9.424
	高山松 P. densata	76.250^b	10.873	14.260
	云南松 P. yunnanensis	67.714^a	9.087	13.420
2012年10月的保存率 Survival rate in Oct. 2012 (%)	油松 P. tabuliformis	63.900^c	9.769	15.288
	高山松 P. densata	18.125^b	30.381	167.619
	云南松 P. yunnanensis	0.143^a	0.378	264.336
2011年11月的封顶率 Rate of bud-set in Nov. 2011 (%)	油松 P. tabuliformis	91.425^c	4.033	4.411
	高山松 P. densata	70.906^b	14.282	20.142
	云南松 P. yunnanensis	10.928^a	20.478	187.390
2012年10月的封顶率 Rate of bud-set in Oct. 2012 (%)	油松 P. tabuliformis	81.900^c	6.976	8.518
	高山松 P. densata	19.834^b	35.432	178.643
	云南松 P. yunnanensis	0.000^a	0.000	-
黄苗比例 Proportion of yellow seedlings (%)	油松 P. tabuliformis	2.300^a	6.845	297.609
	高山松 P. densata	0.406^a	0.481	118.473
	云南松 P. yunnanensis	43.143^b	37.396	86.679
绿苗比例 Proportion of green seedlings (%)	油松 P. tabuliformis	54.350^c	12.849	23.641
	高山松 P. densata	20.688^b	28.561	138.056
	云南松 P. yunnanensis	8.500^a	10.598	124.682
紫苗比例 Proportion of purple seedlings (%)	油松 P. tabuliformis	43.350^a	13.838	31.922
	高山松 P. densata	78.844^b	28.625	36.306
	云南松 P. yunnanensis	47.429^a	34.942	73.672
苗高 Seedling height (cm)	油松 P. tabuliformis	8.286^a	1.242	14.989
	高山松 P. densata	7.763^b	1.070	13.783
	云南松 P. yunnanensis	6.363^c	1.610	25.303
苗冠高 Crown height of seedling (cm)	油松 P. tabuliformis	6.807^a	1.351	19.847
	高山松 P. densata	6.558^b	0.987	15.050
	云南松 P. yunnanensis	5.140^c	1.493	29.047
地径 Ground diameter (mm)	油松 P. tabuliformis	2.486^a	0.124	4.988
	高山松 P. densata	2.093^b	0.270	12.900
	云南松 P. yunnanensis	2.501^a	0.348	13.914

性状 Characters	均方(自由度) MS ( df )				方差分量 Component of variance (%)
性状 Characters	区组间 Among blocks	种间 Among species	群体内 Within population	残差项 Residual	区组间 Among blocks	种间 Among species	群体内 Within population	残差项 Residual
出苗率 Germination rate (%)	0.035 (3)	0.391 (2)^**	0.034 (22)^*	0.016 (72)	2.290	33.700	13.690	50.320
2011年8月的保存率 Survival rate in Aug. 2011 (%)	0.146 (3)^**	0.139 (2)^**	0.042 (22)	0.025 (72)	13.270	8.040	11.530	67.150
2011年9月的保存率 Survival rate in Sept. 2011 (%)	0.126 (3)^**	0.135 (2)^**	0.041 (22)^*	0.024 (72)	11.630	8.110	12.500	67.760
2011年11月的保存率 Survival rate in Nov. 2011 (%)	0.147 (3)^**	0.131 (2)^**	0.039 (22)	0.024 (72)	13.930	7.920	10.340	67.810
一年生苗越冬保存率 Winter survival rate of one-year old seedlings (%)	0.156 (3)^**	0.220 (2)^**	0.034 (22)	0.023 (72)	14.610	15.500	7.900	61.990
2012年10月的保存率 Survival rate in Oct. 2012 (%)	0.039 (3)^*	3.270 (2)^**	0.134 (22)^**	0.014 (72)	0.640	70.830	19.530	9.000
2011年11月的封顶率 Rate of bud-set in Nov. 2011 (%)	0.057 (3)	4.761 (2)^**	0.083 (22)^*	0.043 (72)	0.290	72.640	5.140	21.930
2012年10月的封顶率 Rate of bud-set in Oct. 2012 (%)	0.015 (3)	6.407 (2) ^**	0.168 (22) ^**	0.008 (72)	0.110	79.600	16.840	3.450
黄苗比例 Proportion of yellow seedlings (%)	0.014 (3)	1.457 (2)^**	0.131 (22)^**	0.039 (72)	0.000	39.700	22.490	38.800
绿苗比例 Proportion of green seedlings (%)	0.063 (3)	1.867 (2)^**	0.144 (22)^**	0.025 (72)	1.410	48.260	27.520	22.810
紫苗比例 Proportion of purple seedlings (%)	0.063 (3)	1.307 (2)^**	0.243 (22)^**	0.026 (72)	1.320	28.360	47.740	22.580
苗高 Seedling height (cm)	141.950 (3)^**	252.240 (2)^**	66.330 (22)^**	53.300 (967)	6.870	7.860	15.990	69.290
苗冠高 Crown height of seedling (cm)	161.720 (3)^**	198.700 (2)^**	64.300 (22)^**	56.070 (967)	7.690	5.930	13.470	72.910
地径 Ground diameter (mm)	11.600 (3)^**	17.500 (2)^**	2.510 (22)^**	2.230 (967)	8.670	9.460	8.010	73.860